1 Tutorial goals and set up

1.1 Objectives

Our main objectives are to handle some of the common data problems encountered with marine fastloc GPS data and diving data, using a narwhal dataset provided by Dr. Marianne Marcoux. Specifically, we aim to address the following:

  • Irregular locations

  • Time gaps

  • Including diving covariates

2 Loading the required packages

First, we’ll setup the workspace with required packages.

library(momentuHMM)
library(dplyr)
library(tidyr)
library(lubridate)
library(adehabitatLT)
library(sf)
library(tmap)
library(terra)
library(units)
library(stringr)
library(diveMove)
library(conicfit)
library(car)
library(mitools)
library(doFuture)
library(corrplot)  # Pearson's correlation matrix using cor()

Please also set working directory to “Day2” of the HMM workshop:

setwd("Day2")

3 Import data and initial data processing

For simplicity, we will also only look at the data for the month of August, 2017.

tracks <- read.csv("data/tracks.csv") %>%
  filter(!is.na(x) & !is.na(y)) %>% # remove missing locations
  mutate(
    time = ymd_hms(time), # define time
    loc_class = factor(loc_class, # define location class factor levels
      levels = c("GPS", 3, 2, 1, 0, "A", "B"))) %>% 
  # remove identical records
  filter(!(time == lag(time) & x == lag(x) & y == lag(y) & loc_class == lag(loc_class)),
  # filter only the month of august
  month(time) == 8, day(time) > 7,  day(time) <= 14)

The classic HMM assumes the observation data is in discrete time and that there is no missing data in the predictor variables.

# Calculate time difference between locations
tracks <- tracks %>%
  mutate(dt = ifelse(ID == lead(ID), # If next data row is same individual
    difftime(lead(time), time, units = "mins"), NA
  )) # calculate time difference
# Look at the frequency of time differences
hist(tracks$dt, 1000, main = NA, xlab = "Time difference (min)")

Next, we’ll convert the data to a spatial dataset using the sf package and plot the data. First, we define the coordinate reference system of the original data (in this case WGS84, which is defined by the EPSG code 4326). Next, we will project the data into NAD83(CSRS) UTM zone 21N (EPSG:2962), which will projected the coordinates in meter units with minimal distortion for this data set.

tracks <- tracks %>%
  st_as_sf(coords = c("x", "y")) %>% # converts to an sf object
  st_set_crs(4326) %>% # define CRS
  st_transform(2962) # reproject data to a UTM

For the first part of this tutorial, we’ll use only the fastloc GPS data so we don’t have to deal with location error.

# filter GPS locations only
tracks_gps <- tracks %>% 
  filter(loc_class=="GPS")

We lose some data, particularly near the end of the tracks, but we will integrate ARGOS locations later in this tutorial.

Now, we can map the data using the tmap package to visualize what it looks like.

# plot GPS
tracks_gps %>% 
  group_by(ID) %>% 
  summarise(do_union = FALSE) %>% 
  st_cast("LINESTRING") %>% 
     tm_shape() +
   tm_lines(col = "ID", palette = "Dark2")

3.1 Selecting a time interval for the HMM

There are two key decisions we must make, the temporal resolution to use, and how to address data gaps. The desired resolution depends predominantly on the biological question you are asking as different behaviours and biological processes occur at different spatial and temporal scales (e.g. seasonal migration, daily movement between foraging and resting grounds, and fine scale foraging decisions). Generally, higher resolution data is preferred as it has more information, however it is possible to have too-high of a resolution wherein information from fine-scale variability drowns out the signal from coarse-scale patterns of interest (e.g., seasonal migration). In this case, we will be linking the movement data with high resolution (75 s) dive data to look at finer-scale behaviours (on the order of a few hours). My rule of thumb, is that you want 3-50 data points per behaviour. For behaviours spanning several hours, that roughly corresponds to a desired resolution between 2 min and 60 min.

Let’s see what resolutions may be possible in the data by looking at the most frequent time gaps.

# identify the most frequent dt
tracks_gps %>% 
  {table(.$dt)} %>% 
  sort(decreasing = TRUE) %>% 
  head()
## 
##  10  11   1   9  12   2 
## 299 109 101  75  72  56
# Visualize time differences
hist(tracks_gps$dt, 1000, main = NA, xlab = "Time difference (min)")

# Zoom in on short intervals
hist(tracks_gps$dt, 1000, main = NA, xlab = "Time difference (min)", xlim = c(0,100))

We see that that the most frequent time gap is 10 min, followed by 11, 1, 9, and 12 min. We also see the majority of the gaps are < 60 min, however some are in excess of 600 min. Although it may be possible to use 1 or 2 min resolution data, we will have to address the data gaps. Frequent and large data gaps can be difficult to handle, especially as the number of missing data points approaches or exceeds the existing data; we really want to avoid this. Let’s examine the potential data structure at different resolutions for the different animals.

We first create a function that can calculate the proportion of missing locations we would have for a given resolution.

# Make function to estimate proportion of missing location 
p_na <- function(t_0, t_max, n_loc, resolution) {
  max_n_loc <- length(seq(t_0, t_max, by = as.difftime(resolution, units = "mins")))
  n_NA <- max_n_loc - n_loc
  n_NA / max_n_loc
}

We can now use this function look at the proportion of NAs we would get with 1, 2, 5, 10, and 20 min resolution.

# summarise tracks
track_summary <-
    tracks_gps %>% st_drop_geometry() %>% 
  # limit to core period with data
  group_by(ID) %>% 
    filter(dt >= 1) %>% # remove duplicate time (just for this stage)
    summarise(n_loc = n(), # number of locations
              p_NA_1m = p_na(first(time), last(time), n_loc, 1),  # 1 min 
              p_NA_2m = p_na(first(time), last(time), n_loc, 2),  # 2 min 
              p_NA_5m = p_na(first(time), last(time), n_loc, 5),  # 5 min 
              p_NA_10m =p_na(first(time), last(time), n_loc, 10),  # 10 min 
              p_NA_20m =p_na(first(time), last(time), n_loc, 20))  # 20 min 
track_summary
## # A tibble: 3 × 7
##   ID      n_loc p_NA_1m p_NA_2m p_NA_5m p_NA_10m p_NA_20m
##   <chr>   <int>   <dbl>   <dbl>   <dbl>    <dbl>    <dbl>
## 1 T172062   485   0.950   0.900   0.751    0.501  0.00206
## 2 T172064   479   0.951   0.902   0.754    0.508  0.0164 
## 3 T172066   491   0.949   0.899   0.747    0.493 -0.0124

Here we see that despite the large number of 1 min time steps, at that resolution, >95 /% of the potential locations are missing. Even at the 10 min interval, > 50% of the locations would be missing. Very large data gaps that contribute to much of the missing locations can be excluded from the analysis, therefore, for this tutorial, I will use a 10 min resolution as a compromise between high-resolution information and good temporal coverage.

3.2 Handling data gaps

There are several ways to deal with data gaps, and I will address four 1. Interpolation (linear or statistical) 2. Nullification 3. Path segmentation

3.2.1 Approach 1. Interpolation

For large datasets with few and small gaps, the simplest approach to use linear interpolation. First, let’s identify the most likely minute we have data.

# which minute has the most data
tracks_gps %>% 
  st_drop_geometry() %>%  # must convert back to data.frame
  group_by(ID) %>% 
  summarise(minute = str_sub(minute(time),-1)) %>% 
  table()
## `summarise()` has grouped output by 'ID'. You can override using the `.groups`
## argument.
##          minute
## ID          0   1   2   3   4   5   6   7   8   9
##   T172062  81  78  62  52  37  40  40  45  37  32
##   T172064  77  68  71  67  32  32  40  32  31  38
##   T172066 138  52  73  48  44  42  37  27  22  18

It looks like for all three tracks, the most amount of locations fall on 0 min (i.e., 10, 20, 30, 40, 50, 60 min). Next, for each track, we will create a vector of times in which to estimate locations.

# convert tracks back to data.frame with xy coordinates
tracks_gps_ls <- tracks_gps %>% 
  mutate(x = st_coordinates(tracks_gps)[,"X"],  
         y = st_coordinates(tracks_gps)[,"Y"]) %>%
  st_drop_geometry() %>% 
    split(.,.$ID)  # split into list

# create full time series on which to estimate locations rounded to the nearest 10 min
tracks_gps_ls_time <- tracks_gps %>% 
  st_drop_geometry() %>%  # convert to data.frame
  group_by(ID) %>%
  summarise(time = seq(round_date(first(time), "10 min"), 
                       round_date(last(time), "10 min"),
                       by = 60*10)) %>% 
  split(.,.$ID)  # split into list
## `summarise()` has grouped output by 'ID'. You can override using the `.groups`
## argument.

Now, we can interpolate the locations for each track.

# function to create a data frame with approximated locations
approx_locs <- function(tracks, times){
  data.frame(ID = times$ID,
        time = times$time,
        x = approx(tracks$time, tracks$x,
                   xout = times$time)$y,
        y = approx(tracks$time, tracks$y,                               
                   xout = times$time)$y)
}

# Interpolate the location at the times from the sequence
tracks_gps_linear <- mapply(approx_locs, tracks_gps_ls, tracks_gps_ls_time,
                          SIMPLIFY = F) %>% 
  do.call("rbind", .)  # convert list of tracks back to a single data.frame

# remove row names added by rbind
rownames(tracks_gps_linear) <- NULL 

# plot locations
plot(tracks_gps_linear$x, tracks_gps_linear$y, pch = 20, col = "red", xlab = "x", ylab = "y", asp = 1)
points(st_coordinates(tracks_gps)[,"X"], st_coordinates(tracks_gps)[,"Y"], pch = 20)

Looks like it works. Let’s try fitting an HMM to this. First, lets prepare the data using prepData and plot the data to estimate starting parameters.

prep_gps_linear <- prepData(tracks_gps_linear, type = "UTM")

plot(prep_gps_linear, ask = F)

# Setting up the starting values
mu0 <- c(50, 500) # Mean step length
sigma0 <- c(50, 500) # Sd of the step length
kappa0 <- c(0.1, 1) # Turning angle concentration parameter (kappa > 0)

Ok, were are ready. Let’s fit the HMM

set.seed(1)
# Fit a 2 state HMM
HMM_gps_linear <- fitHMM(prep_gps_linear, nbState = 2, dist = list(step = "gamma",
angle = "vm"), Par0 = list(step = c(mu0, sigma0), angle = kappa0), formula = ~ 1)

plot(HMM_gps_linear, ask = FALSE)
## Decoding state sequence... DONE

That model does not look good. It’s clear from the mapped states that state 1 largely represents the original data, while state 2 primarily represents the interpolated data. This is a common problem when data gaps are frequent or large such that the information in the interpolated data outweighs the signal from the original observations. Generally, I would only use linear interpolation when data gaps are small (< 3 locations) or relatively infrequent (< 20 % of the modelled locations). In our data, some gaps are > 5 hours (30 locations) and > 50% of the modelled locations are interpolated. So we need to use another approach.

A slightly better way to interpolate locations is to fit a correlated random walk (CRW) model to the data and predict the most likely locations. momentuHMM contains wrapper functions to interpolate missing locations by fitting continuous-time CRW to the data based on the crawl package by Devin Johnson and Josh London. There are many options to fit the crawl model, and a detailed tutorial for analysis with crawl is available here: https://jmlondon.github.io/crawl-workshop/crawl-practical.html. Let’s try to fit the most basic model using the wrapper function crawlWrap.

set.seed(1) # crawl often fails, so I recommend always setting a seed 
# fit crawl
crw_gps_10 <- crawlWrap(obsData = tracks_gps, timeStep = "10 mins")
## Fitting 3 track(s) using crawl::crwMLE...
## crawl 2.2.1 (2018-09-12) 
##  Demos and documentation can be found at our new GitHub repository:
##  https://dsjohnson.github.io/crawl_examples/
## Loading required package: rngtools
## Individual T172062...
## Beginning SANN initialization ...
## Beginning likelihood optimization ...
## Individual T172064...
## Beginning SANN initialization ...
## Beginning likelihood optimization ...
## Individual T172066...
## Beginning SANN initialization ...
## Beginning likelihood optimization ...
## Warning in sqrt(diag(Cmat)): NaNs produced
## DONE
## Warning in eval(c.expr, envir = args, enclos = envir): crawl::crwMLE for
## individual T172066 has NaN variance estimate(s)
## Predicting locations (and uncertainty) at 10 mins time steps for 3 track(s) using crawl::crwPredict...
## Warning in nax != nay: longer object length is not a multiple of shorter object
## length
## Warning in nax != nay: longer object length is not a multiple of shorter object
## length

## Warning in nax != nay: longer object length is not a multiple of shorter object
## length
## DONE
# plot fit tracks
plot(crw_gps_10, ask = FALSE)

Notice how the predicted tracks do not make perfectly straight lines through missing sections (particularly noticeable in T172062). Next, we will extract the predicted locations and add them to the observed data.

# filter predicted locations
tracks_gps_crw <- data.frame(crw_gps_10$crwPredict) %>% 
  filter(locType == "p") %>% 
  dplyr::select(mu.x, mu.y, time,
         ID) %>% 
  dplyr::rename(x = "mu.x", y = "mu.y")

Now, let’s try to fit the same HMM as before on this data using the same starting parameters.

set.seed(1)
# prep data 
prep_gps_crw <- prepData(tracks_gps_crw, type = "UTM")

# Setting up the starting values
mu0 <- c(50, 500) # Mean step length
sigma0 <- c(50, 500) # Sd of the step length
kappa0 <- c(0.1, 1) # Turning angle concentration parameter (kappa > 0)

# Fit a 2 state HMM
HMM_gps_crw <- fitHMM(prep_gps_crw, nbState = 2, dist = list(step = "gamma",
    angle = "vm"), Par0 = list(step = c(mu0, sigma0), angle = kappa0))

plot(HMM_gps_crw, ask = F)
## Decoding state sequence... DONE

That’s looking much better. It looks like state 1 represents a low-speed, high tortuosity resident state, while state 2 represents higher-speed, low tortuosity travelling state.

In many instances, this model may be sufficient. However, the significant proportion of interpolated locations used to fit the model is likely to affect our results. For example, the large interpolated gaps are still relatively straightened out and a very consistent speed, and may skew the definition of state 2 in particular. Simply interpreting the results with this issue in mind can be adequate when we only have few interpolated locations. Below, we will explore more formal ways to address this problem, which can be particularly useful when we have very large numbers of interpolated locations.

3.2.2 Approach 2. Nullify data gaps

One strategy to address large data gaps is to nullify the data streams (i.e., step length and turning angle) during moderate/large interpolated gaps where we expect that the estimated movement has is largely independent of the observed data before or after the gap. The maximum size of a gap to allow depends on the frequency of the missing data, frequency of locations, study species, and behaviours of interest. In this case, I will nullify predicted locations in gaps larger than 60 min. First, we will identify which steps need to be nullified, then we will prepare the data and nullify the estimated step and angle data streams. We will do this again later in the tutorial, so we will wrap it into a function called prepData_NAGaps.

# define function to replace step and turn angle of large gaps with NA
NA_gaps <- function(tracks, times){
  # rows where location is within a large gap
  rows <- which(
    rowSums(apply(times, 1, function(X, tracks){
      dplyr::between(tracks, 
                     as.POSIXct(X[1], tz = "UTC"),
                     as.POSIXct(X[2], tz = "UTC"))
    }, tracks$time))==1)
  tracks$step[rows] <- NA
  tracks$angle[rows] <- NA
  return(tracks)
}

# define function to identify and nullify gaps
prepData_NAGaps <- function(track_list, tracks_crw, res, max_gap, ...){
  # for each ID, identify which rows have gaps >= max_gap 
  gaps_ls_rows <- lapply(track_list, function(x){
    which(difftime(lead(x$time), x$time, units = "mins") >= max_gap)
  })
  
  # create sequence of times for each track from gaps >= 60 min
  gap_ls <- mapply(FUN = function(track, gaps){
    # identify start and end date of each gap
    gaps_ls_srt_end <- list(start = ceiling_date(track$time[gaps], paste(res, "min")),
                            end = floor_date(track$time[gaps+1], paste(res, "min")))
    # combine into single vector for each track
    data.frame(start = gaps_ls_srt_end$start, end = gaps_ls_srt_end$end)
  },
  track_list, gaps_ls_rows, SIMPLIFY = F)
  
  # prep data and list by ID
  prep_tracks <- prepData(tracks_crw, ...) %>% 
    {split(., .$ID)}
  
  # Interpolate the location at the times from the sequence
  mapply(FUN = NA_gaps, prep_tracks, gap_ls,
         SIMPLIFY = F) %>% 
    do.call("rbind", .) # convert list of tracks back to a single data.frame
}
  
prep_tracks_gps_crw_NAgaps <- prepData_NAGaps(tracks_gps_ls, tracks_gps_crw, 10, 60, type = "UTM")

Now, let’s try to fit the same HMM as above to this data with large gaps nullified.

set.seed(1)
# Setting up the starting values
mu0 <- c(50, 500) # Mean step length
sigma0 <- c(50, 500) # Sd of the step length
kappa0 <- c(0.1, 1) # Turning angle concentration parameter (kappa > 0)

# Fit a 2 state HMM
HMM_gps_crw_NAgaps <- fitHMM(prep_tracks_gps_crw_NAgaps, nbState = 2, dist = list(step = "gamma", angle = "vm"), Par0 = list(step = c(mu0, sigma0), angle = kappa0))

plot(HMM_gps_crw_NAgaps, ask = FALSE)
## Decoding state sequence... DONE

Visually, the difference is subtle.

HMM_gps_crw$mle[c("step", "angle")]
## $step
##       state 1  state 2
## mean 258.1724 705.2750
## sd   158.3769 255.9088
## 
## $angle
##                state 1  state 2
## mean          0.000000  0.00000
## concentration 2.706698 23.60863
HMM_gps_crw_NAgaps$mle[c("step", "angle")]
## $step
##       state 1  state 2
## mean 282.3941 757.6624
## sd   156.3310 248.9599
## 
## $angle
##                state 1  state 2
## mean          0.000000  0.00000
## concentration 2.476524 19.03249

However, the estimated parameters are quite different whether you account for the large gaps in data. When you nullify large gaps, the mean step length for both states is higher, and the turn angle concentration parameters is lower for both states (i.e., more tortuous). The fact that the parameters change for both states, suggests that the large gaps skewed the parameterisation of both states.

3.2.3 Approach 3. Path segmentation

Another strategy to deal with larger gaps is to segment the tracks with a new individual ID. This may be particularly appropriate for gaps where we may reasonably expect that the the underlying states are effectively independent of one another. Specifically, we may ask, over what period of time does the behaviour of the animal affect the subsequent behaviour. In this case, we may expect that the behaviour of a narwhal depends on the behaviour over the proceeding several hours, however is independent after 24 hours. We can split the tracks for gaps larger than a predetermined threshold by iterating the ID column. We will not implement this approach in this tutorial, however, it can be done using the following code:

gap_thresh <- 3*60 # in hours (3h for illustration)
# gaps no more than 6h per segment
new_ID <- (tracks_gps$dt > gap_thresh | tracks_gps$ID != lag(tracks_gps$ID)) %>%  # if dif.time > gap_thresh or new ID
  replace_na(TRUE) %>%  # replace first NA with ID = 1
  cumsum() %>%  # iterate ID 
  paste(tracks_gps$ID, ., sep = "_")
# then smaller gaps <= gap_thresh can be interpolated with crawlWrap

3.3 Integrating dive data

Let’s import the dive data and explore it.

dives <- read.csv("data/dives.csv") %>% 
  mutate(time = ymd_hms(time)) %>% 
  filter(month(time) == 8, day(time) > 7,  day(time) <= 14)

head(dives)
##        ID                time depth   dt
## 1 T172062 2017-08-08 00:00:00   1.0 1.25
## 2 T172062 2017-08-08 00:01:15   7.5 1.25
## 3 T172062 2017-08-08 00:02:30  11.0 1.25
## 4 T172062 2017-08-08 00:03:45   5.5 1.25
## 5 T172062 2017-08-08 00:05:00   3.5 1.25
## 6 T172062 2017-08-08 00:06:15   1.0 1.25
table(dives$dt)
## 
##   1.25  61.25 121.25 181.25 241.25 301.25 361.25 
##  18547     52     16      6      1      1      1

It appears as though there are relatively few gaps, however the gaps that exist are relatively long (> 60 mins). Therefore, I suggest that we should regularize and nullify the dive data.

# regularise data 
dives <- dives %>% 
  group_by(ID) %>%
  # regularise time series by 1.25 min
  summarise(time = seq(first(time), last(time), by = 1.25*60)) %>%
  # merge regularised time with original dive
  left_join(dives, by = c("ID", "time"))
## `summarise()` has grouped output by 'ID'. You can override using the `.groups`
## argument.

Next, we have to summarise the time series into concrete data streams that can be modelled in the HMM. We can use the diveMove package to identify individual dives and calculate dive statistics. We must first convert the depth data to the TDR class for each whale. As we are working with multiple whales, we will use lapply to apply the createTDR to each whale – we will also use lapply for most of diveMove functions.

dives_ls <- split(dives, dives$ID) 
dive_TDR <- lapply(dives_ls, function(data){
  createTDR(time = data$time, depth = data$depth, dtime = 1.25*60, file = "data/dives.csv")
  })

# generate interactive plot
plotTDR(dive_TDR[[1]]) # note: try zooming in to part of the dive

Next, we must use the calibrateDepth to calibrate the depth data. This function identifies wet and dry periods (for animals that haul out), applies a zero-offset correction (ZOC), and identifies all dives in the record and their phases. ZOC can be done using either the offset or filter method. In this case, we will assume the depth data is accurate and does not require an offset. We should also specify dive.thr, which represents the threshold depth below which an underwater phase should be considered a dive – in this case, we will set it at 8 m. There are many other optional parameters to identify dives that we will not get into in this tutorial (See vignette("diveMove") for details).

# calibrate TDR and identify dive phases
dive_TDR_calib <- lapply(X = dive_TDR, FUN = calibrateDepth, zoc.method = "offset", offset = 0, dive.thr = 8)
## Record is truncated at the beginning and at the end
## 39 phases detected
## 818 dives detected
## Record is truncated at the beginning and at the end
## 75 phases detected
## 484 dives detected
## Record is truncated at the beginning and at the end
## 41 phases detected
## 628 dives detected
# interactive plot of calibrated DTR
plotTDR(dive_TDR_calib[[1]], surface = TRUE)

In the interactive plot, try zooming into one of the dives and hover mouse over plot to preview the phases identified by `calibrateDepth’. The phases identified correspond to descent (D), descent/bottom (DB), bottom (B), bottom/ascent (BA), ascent (A), and surface (X). We can plot a specific dives as follows:

plotTDR(dive_TDR_calib[[1]], diveNo = 1:300, surface = TRUE)

Now, we can calculate summary statistics for each dive using the function diveStats. There are many dive metrics that are estimated, and which ones to retain are species, data, and question specific. In this case, we will retain 8 from those calculated by diveStats: dive time, bottom time, maximum depth, bottom distance (measure of “wiggling while at the bottom), post-dive duration.

# calculate dive stats and add dive.id to each dive 
dive_sum <- lapply(dive_TDR_calib, function(data){
  mutate(diveStats(data, depth.deriv = F), dive.id = row_number()) %>% 
    dplyr::select(dive.id, divetim, botttim, maxdep, bottdist, postdive.dur)}) # select variables of interest

# add dive.id with depth data to each depth record
dives_ls <- mapply(function(TDR, dives){
  mutate(TDR, dive.id = dives@dive.activity$dive.id)
  }, TDR = dives_ls, dives = dive_TDR_calib, SIMPLIFY = F)

# join TDR data with dive summary data
dives_ls <- mapply(function(TDR, dive_sum){
  left_join(TDR, dive_sum, by = "dive.id")
  }, TDR = dives_ls, dive_sum = dive_sum, SIMPLIFY = F)

# convert dive_ls back to a data.frame
dives_df <- do.call(rbind, dives_ls)

Next, we will replace any bottom time of a valid dive (dive.id > 0) to 75 s, since at least some time must be spent at the bottom. Then, we will calculate one additional metric as a proxy for dive shape; the ratio of bottom time to dive time. Dives where \(\leq20\%\) of the time is spent at the bottom represent V-shaped dives, U-shaped dives are represented when \(>20\) and \(\leq50\%\) is spent at the bottom, and square dives are represented when \(>50\%\) of the time is spent at the bottom.

# replace NA bottom time of valid dives
dives_df$botttim[dives_df$dive.id > 0 & is.na(dives_df$botttim)] <- 75
# calculate proportion time at bottom
dives_df <- dives_df %>% 
  mutate(propbott = botttim/divetim)
# remove "dives" with no duration
dives_df <- dives_df %>% 
  filter(!(dive.id > 0 & is.na(divetim)))

The next issue is that the dive data is at a different temporal resolution (75 s) than the location data (10 min). There are two options to include both data streams in the same HMM. First, we can choose to implement a hierarchical HMM, however, this is more complicated, and will be covered in tomorrow’s tutorial. The second, which we will use here, is to summarise the depth/dive data to a 10 min resolution and include them as additional data streams with step length and turning angle.

dives_sum <- dives_df %>% 
  # round time to same interval as location data
  mutate(time = floor_date(time, "10 min")) %>% 
  # group rows by time interval
  group_by(ID, time) %>% 
  # summarise data
  summarise(NA_t          = sum(is.na(depth))*1.25,
            surf_t        = sum(dive.id == 0)*1.25,
            mean_dep      = ifelse(NA_t == 10, NA, mean(na.rm = T, depth)),
            max_dep       = ifelse(NA_t == 10, NA, max(na.rm = T, depth)),
            dive_t        = ifelse(NA_t == 10, NA, sum(na.rm = T, divetim)),
            bott_t        = ifelse(dive_t < 5, NA, sum(na.rm = T, botttim)),
            prop_bott     = ifelse(dive_t < 5, NA, max(na.rm = T, propbott)),
            max_dive_dep  = ifelse(dive_t < 5, NA, max(na.rm = T, maxdep)),
            bott_dist     = ifelse(dive_t < 5, NA, max(na.rm = T, bottdist)),
            post_dive_dur = ifelse(dive_t < 5, NA, max(na.rm = T, postdive.dur))) %>% 
  # remove -Inf values (typically error)
  filter(!is.infinite(dive_t) & !is.infinite(bott_dist))
## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf

## Warning in max(na.rm = T, bottdist): no non-missing arguments to max; returning
## -Inf
## `summarise()` has grouped output by 'ID'. You can override using the `.groups`
## argument.
# preview
head(dives_sum)
## # A tibble: 6 × 12
## # Groups:   ID [1]
##   ID      time                 NA_t surf_t mean_dep max_dep dive_t bott_t
##   <chr>   <dttm>              <dbl>  <dbl>    <dbl>   <dbl>  <dbl>  <dbl>
## 1 T172062 2017-08-08 00:10:00     0   3.75    17.3     31     1875    375
## 2 T172062 2017-08-08 00:20:00     0   6.25     7.56    20     1875   1125
## 3 T172062 2017-08-08 00:30:00     0   0      109.     123     7200   4200
## 4 T172062 2017-08-08 00:40:00     0   5       20.9     88     2100   1200
## 5 T172062 2017-08-08 00:50:00     0   5        9.94    24.5   1275    450
## 6 T172062 2017-08-08 01:00:00     0   1.25    53.2     76.5   4200   1575
## # … with 4 more variables: prop_bott <dbl>, max_dive_dep <dbl>,
## #   bott_dist <dbl>, post_dive_dur <dbl>

Note, we used 2 different ifelse statements. First, ifelse(NA_t == 10, ...) ensured that we only calculate mean and maximum depth for periods where we have at least 1.25 min of depth data. Second, ifelse(surf_t > 5, ...) ensured that we only calculate dive metrics when at least half of the 10 min interval is spent in a dive, otherwise the animal is assumed to be at the surface and its dive metric is NA.

Any of the eight variables can be used as data streams in the HMM, however, including too many would significantly increase the number of parameters to estimate, and consequently computation time. Therefore, we must select which variables to use, which there are different several approaches: 1. Variable can be selected using expert on the species behaviour and research question. 2. We preferably want to avoid variable with a lot of missing data. 3. The data streams should exhibit variation and evidence of clustering or multi-modality that may be tied to the underlying behaviours. 4. Data streams with no variation or conversely are very noisy contain little information on underlying behaviour. 5. We want to avoid variables that are overdispersed and which would be difficult to describe using a statistical distribution. 6. Given that they are tied to autocorrelated behaviours, they too should exhibit some – but not too much – autocorrelation. 7. One of HMM’s key assumptions that the data stream are be independent of each other, therefore, we should avoid select highly co-linear data streams.

Assuming all the variables are biologically relevant, let’s look at structure in the data. I will also add a subjective score based on the results (first number after # symbol).

vars <- c("mean_dep","max_dep","dive_t","bott_t","prop_bott",
          "max_dive_dep","bott_dist","post_dive_dur")
# First, missing data
dives_sum %>% 
  summarise(mean_dep      = sum(is.na(mean_dep)),      # 2
            max_dep       = sum(is.na(max_dep)),       # 2
            dive_t        = sum(is.na(dive_t)),        # 2
            bott_t        = sum(is.na(bott_t)),        # 1
            prop_bott     = sum(is.na(prop_bott)),     # 1
            max_dive_dep  = sum(is.na(max_dive_dep)),  # 1
            bott_dist     = sum(is.na(bott_dist)),     # 1
            post_dive_dur = sum(is.na(post_dive_dur))) # 1
## # A tibble: 3 × 9
##   ID      mean_dep max_dep dive_t bott_t prop_bott max_dive_dep bott_dist
##   <chr>      <int>   <int>  <int>  <int>     <int>        <int>     <int>
## 1 T172062      120     120    120    216       216          216       216
## 2 T172064      366     366    366    476       476          476       476
## 3 T172066      210     210    210    359       359          359       359
## # … with 1 more variable: post_dive_dur <int>
# Second, evidence of multi-modality, balanced variation, and no over-dispersion
# as the dive variables are zero-bound, applying a log transformation makes it easier to see structure in the distirbution 
# dives_sum %>% 
#   ungroup() %>% 
#   dplyr::select(vars) %>% 
#   apply(2, hist, 100)
hist(log(dives_sum$mean_dep), 100)      # 2 good structure, high dispersion

hist(log(dives_sum$max_dep), 100)       # 3 great structure, moderate dispersion

hist(log(dives_sum$dive_t), 100)        # 1 little structure, fragmented distribution

hist(log(dives_sum$bott_t), 100)        # 0 little structure, fragmented distribution

hist(logit(dives_sum$prop_bott), 100)   # 0 no structure, fragmented distribution
## Warning in logit(dives_sum$prop_bott): proportions remapped to (0.025, 0.975)

hist(log(dives_sum$max_dive_dep), 100)  # 2 good structure, low dispersion

hist(log(dives_sum$bott_dist), 100)     # 2 moderate structure, high dispersion

hist(log(dives_sum$post_dive_dur), 100) # 0 little structure, high dispersion

# Third, balanced autocorrelation
dives_sum_filter <- filter(dives_sum, ID == "T172062")
acf(dives_sum_filter$mean_dep, na.action = na.pass)       # 1 ACF~8 some variability

acf(dives_sum_filter$max_dep, na.action = na.pass)        # 2 ACF~8 gradual decline

acf(dives_sum_filter$dive_t, na.action = na.pass)         # 2 ACF~7 gradual decline

acf(dives_sum_filter$bott_t, na.action = na.pass)         # 0 ACF~3

acf(dives_sum_filter$prop_bott, na.action = na.pass)      # 0 ACF~2

acf(dives_sum_filter$max_dive_dep, na.action = na.pass)   # 1 gradual but high ACF~22

acf(dives_sum_filter$bott_dist, na.action = na.pass)      # 1 low ACF~4

acf(dives_sum_filter$post_dive_dur, na.action = na.pass)  # 0 low ACF~1

# subjective score
data.frame(var = c("mean_dep", "max_dep", "dive_t", "bott_t", "prop_bott", 
                   "max_dive_dep", "bott_dist", "post_dive_dur"),
           score = c(5, 6, 5, 1, 
                     1, 4, 4, 1))
##             var score
## 1      mean_dep     5
## 2       max_dep     6
## 3        dive_t     5
## 4        bott_t     1
## 5     prop_bott     1
## 6  max_dive_dep     4
## 7     bott_dist     4
## 8 post_dive_dur     1

From my subjective interpretation of these outputs, I think the five most promising variables to include are maximum depth, dive time, mean depth, maximum dive depth, and bottom distribution. Next, let’s merge the dive data streams the location data.

tracks_dives <- left_join(prep_tracks_gps_crw_NAgaps, 
                          dives_sum[,c("ID", "time", "max_dep", "dive_t", 
                                       "mean_dep", "max_dive_dep", "bott_dist")], 
                          by = c("ID", "time"))

Now, we must check for co-linearity between the data streams to select 1–3 to use in the HMM. We can check co-linearity using the Pearson’s correlation matrix.

# calculate and plot check Paerson's correlation matrix
tracks_dives[,c("max_dep", "dive_t", "mean_dep", "max_dive_dep", 
             "bott_dist", "step", "angle")] %>% 
  na.omit() %>%
  cor() %>% 
  corrplot(method="number")

step and angle seem quit independent from all variables as does bott_dist. Unfortunately, there is quite high correlation between the first four dive data streams, se we will have to select which to use. dive_t and mean_dep have the lowest correlation with bott_dist and step, however max_dep had the highest subjective score. For the purposes of this tutorial, I will use max_dep and bott_dist.

To get starting parameters, we can fit an HMM to each one independently. We will use the gamma distribution for both and for now will use the same starting parameters.

# identify whether there is 0 data
tracks_dives %>% 
  summarise(max_dep = sum(max_dep == 0, na.rm = T), 
            bott_dist = sum(bott_dist == 0, na.rm = T)) 
##   max_dep bott_dist
## 1       0       628
# therefore, we need to include zero-mass parameters for bott_dist

# starting parameters (will use same ones for both for now)
mu0 <- c(130, 180)  # mean
sigma0 <- c(60, 90)  # sd
zm <- c(0.1, 0.1)  # zero mass, where applicable

# fit dive-only HMMs
set.seed(1)
HMM_max_dep <- fitHMM(tracks_dives, nbStates = 2, dist = list(max_dep = "gamma"), 
                      Par0 = list(max_dep = c(mu0, sigma0)))
HMM_bott_dist <- fitHMM(tracks_dives, nbStates = 2, dist = list(bott_dist = "gamma"), 
                        Par0 = list(bott_dist = c(mu0, sigma0, zm)))

Next, we can integrate these into one HMM together with step length and turning angle. When we specify the starting parameters, we want to think about how the states may look. For example, we might expect one resident state with slower movement, lower angular concentration, deeper dives, and greater at-depth variability. The second state may be travel, with faster movement, greater angular concentration, shallower dives, and less at-depth variability.

# prep model
stateNames = c("resident", "travel")
nbStates = length(stateNames)
dist = list(step = "gamma", angle = "vm", max_dep = "gamma", bott_dist =  "gamma")

# Starting Pars 
  # view Pars from previous HMMs
  getPar(HMM_gps_crw_NAgaps)$Par  # state 1 ~ resident, state 2 ~ travel
## $step
## [1] 282.3941 757.6624 156.3310 248.9599
## 
## $angle
## [1]  2.476524 19.032493
  getPar(HMM_max_dep)$Par  # state 1 ~ travel, state 2 ~ resident
## $max_dep
## [1]  30.15317 407.61829  29.05073 163.55047
  getPar(HMM_bott_dist)$Par  # state 1 ~ travel, state 2 ~ resident
## $bott_dist
## [1]  23.9091351 315.3866789  24.7190593 203.5173936   0.2223528   0.5215131
  # therefore must switch parameter estimates in HMM_max_dep & HMM_bott_dist
  
  # combine starting Pars 
  step0 <- getPar(HMM_gps_crw_NAgaps)$Par$step
  angle0 <- getPar(HMM_gps_crw_NAgaps)$Par$angle
  max_dep0 <- c(getPar(HMM_max_dep)$Par$max_dep[c(2,1)],  # mu1, mu2
                getPar(HMM_max_dep)$Par$max_dep[c(4,3)])  # sd1, sd2
  bott_dist0 <- c(getPar(HMM_bott_dist)$Par$bott_dist[c(2,1)],  # mu1, mu2
                  getPar(HMM_bott_dist)$Par$bott_dist[c(4,3)],  # sd1, sd2
                  getPar(HMM_bott_dist)$Par$bott_dist[c(6,5)])  # zm1, zm2
  Par0 <- list(step = step0, angle = angle0, max_dep = max_dep0, bott_dist = bott_dist0)

Now, we can fit the HMM with movement and dive variables.

set.seed(1)
HMM_move_dive <- fitHMM(tracks_dives, nbStates=nbStates, stateNames=stateNames, dist=dist, Par0=Par0)

Let’s see what it looks like.

plot(HMM_move_dive, breaks = 100, ask = F)
## Decoding state sequence... DONE

plotPR(HMM_move_dive)
## Computing pseudo-residuals...

The tracks look interesting. But there are some issues in the pseudo residuals.

Looking at the QQ plots the model appears to: - overestimate the number of fast movement steps - underestimating higher turning angles - underestimate shallow dives and over estimate deep dives - really wonky description of bott_dist The autocorrelation functions suggest there is remnant autocorrelation in step and max_dep that are not well described by the model. Together, this information suggests that there may be additional states that are not represented. Let’s try to add one more state with an intermediate speed, lower angle concentration, and shallow dives.

# define states
stateNames <- c("resident", "travel", "search")
nbStates <- length(stateNames)

# Starting Pars 
  # get Pars from last HMM_move_dive HMM
  Pars <- getPar(HMM_move_dive)$Par 

  # combine starting Pars 
  step0 <- c(Pars$step[c(1,2)], mean(Pars$step[c(1,2)]),  # mu
             Pars$step[c(3,4)], mean(Pars$step[c(3,4)]))  # sd
  angle0 <- c(Pars$angle, 3)
  max_dep0 <- c(Pars$max_dep[c(1,2)], 25, # mu
                Pars$max_dep[c(3,4)], 10) # sd
  bott_dist0 <- c(Pars$bott_dist[c(1,2)], mean(Pars$bott_dist[c(1,2)]), # mu
                  Pars$bott_dist[c(3,4)], mean(Pars$bott_dist[c(3,4)]), # sd
                  Pars$bott_dist[c(5,6)], mean(Pars$bott_dist[c(5,6)]))# zm
  Par0 <- list(step = step0, angle = angle0, max_dep = max_dep0, bott_dist = bott_dist0)

# fit 3-state HMM
set.seed(1)
HMM_move_dive_3s <- fitHMM(tracks_dives, nbStates=nbStates, stateNames=stateNames, dist=dist, Par0=Par0)
plotPR(HMM_move_dive_3s)
## Computing pseudo-residuals...

plot(HMM_move_dive_3s, breaks = 100, ask = FALSE)
## Decoding state sequence... DONE

Interestingly, the step and angle QQ-plots were not improved much, though the step ACF was improved. However, compared to the 2-state model, there was a drastic improvement in QQ-plot and ACF the max_dep data stream and marginal improvement in the bott_dist data stream. The newly described state appears to have very low step length, really wide turning angle, minimal diving, and minimal at-depth activity, which may actually be indicative of resting. The “resident” state has intermediate speed and turning angle, but very deep dives and high at-depth activity, suggesting it may represent foraging. Let’s try to fit one more 4-state HMM, to try and address the remaining residuals: intermediate speed, lower angle concentration, and more intermediate dives, which may represent searching behaviour.

# define states
stateNames <- c("forage", "travel", "rest", "search")
nbStates <- length(stateNames)
# Starting Pars 
  # get Pars from last HMM_move_dive HMM
  Pars <- getPar(HMM_move_dive_3s)$Par 

  # combine starting Pars 
  step0 <- c(Pars$step[c(1:3)], mean(Pars$step[3]),  # mu
             Pars$step[c(4:6)], mean(Pars$step[6]))  # sd
  angle0 <- c(Pars$angle, Pars$angle[3])
  max_dep0 <- c(Pars$max_dep[c(1:2)], Pars$max_dep[3]/2, Pars$max_dep[3], # mu
                Pars$max_dep[c(4:5)], Pars$max_dep[6]/2, Pars$max_dep[6]) # sd
  bott_dist0 <- c(Pars$bott_dist[c(1:3)], Pars$bott_dist[3], # mu
                  Pars$bott_dist[c(4:6)], Pars$bott_dist[6], # sd
                  Pars$bott_dist[c(7:9)], Pars$bott_dist[9]) # zm
  Par0 <- list(step = step0, angle = angle0, max_dep = max_dep0, bott_dist = bott_dist0)

# fit 4-state HM
set.seed(1)
HMM_move_dive_4s <- fitHMM(tracks_dives, nbStates=nbStates, stateNames=stateNames, dist=dist, Par0=Par0)
## =======================================================================
## Fitting HMM with 4 states and 4 data streams
## -----------------------------------------------------------------------
##  step ~ gamma(mean=~1, sd=~1)
##  angle ~ vm(concentration=~1)
##  max_dep ~ gamma(mean=~1, sd=~1)
##  bott_dist ~ gamma(mean=~1, sd=~1, zeromass=~1)
## 
##  Transition probability matrix formula: ~1
## 
##  Initial distribution formula: ~1
## =======================================================================
## DONE
# view output
plotPR(HMM_move_dive_4s)
## Computing pseudo-residuals...

plot(HMM_move_dive_4s, breaks = 200)
## Decoding state sequence... DONE

The QQ-plot for step is quite a bit improved, as is for bott_dist. The ACF for step is also quite improved. At this point, I’m not sure the model can be significantly improved with the existing data. Although HMMs with different number of states generally shouldn’t be compared, since AIC generally favours more states, it is a good sanity check.

AIC(HMM_move_dive, HMM_move_dive_3s, HMM_move_dive_4s)
##              Model      AIC
## 1 HMM_move_dive_4s 70831.17
## 2 HMM_move_dive_3s 71706.75
## 3    HMM_move_dive 73241.60

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